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Filogenia E Evolução De Ouriços-Cacheiros (Rodentia

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Filogenia E Evolução De Ouriços-Cacheiros (Rodentia UNIVERSIDADE FEDERAL DO ESPÍRITO SANTO CENTRO DE CIÊNCIAS HUMANAS E NATURAIS PROGRAMA DE PÓS-GRADUAÇÃO EM CIÊNCIAS BIOLÓGICAS Filogenia e evolução de ouriços-cacheiros (Rodentia: Erethizontidae) Vilacio Caldara Junior Vitória, ES Fevereiro, 2013 ii UNIVERSIDADE FEDERAL DO ESPÍRITO SANTO CENTRO DE CIÊNCIAS HUMANAS E NATURAIS PROGRAMA DE PÓS-GRADUAÇÃO EM CIÊNCIAS BIOLÓGICAS Filogenia e evolução de ouriços-cacheiros (Rodentia: Erethizontidae) Vilacio Caldara Junior Orientador: Yuri Luiz Reis Leite Tese submetida ao Programa de Pós-graduação em Ciências Biológicas (Biologia Animal) da Universidade Federal do Espírito Santo como requisito parcial para a obtenção do grau de Doutor em Biologia Animal Vitória, ES Fevereiro, 2013 "A ignorância gera confiança com mais frequência do que o conhecimento: são aqueles que sabem pouco, e não aqueles que sabem muito, que tão positivamente afirmam que esse ou aquele problema jamais será resolvido pela ciência." Charles Darwin ii Agradecimentos Ao meu orientador Yuri Luiz Reis Leite, por mais essa oportunidade. Agradeço ainda pelo apoio financeiro e operacional e, principalmente, pela grande contribuição intelectual para a concepção e execução deste trabalho. À minha "marida" Sílvia Ramira Lopes Caldara, por todo carinho, dedicação e incentivo. Agradeço pelo auxílio nos trabalhos laboratoriais no Núcleo de Genética Aplicada à Conservação da Biodiversidade (NGACB-UFES) e na análise e interpretação de dados. Aos meus pais Vilacio Caldara e Maria Arevabene Caldara e aos meus irmãos Luciana, Lucimar e Adauto, pelo carinho, afeto e compreensão. Aos professores do Programa de Pós-Graduação em Ciências Biológicas da UFES (PPGBAN), por novamente terem concedido grande volume de conhecimentos para meu crescimento como cientista. Em especial à Leonora Pires Costa, Albert David Ditchfield e Celso Oliveira Azevedo pelas excelentes e indispensáveis discussões que realizamos durante meu processo de qualificação. Ao Ariel Sessa, secretário executivo do PPGBAN, pela ajuda para a obtenção de recursos e documentos. À Juliana de Freitas Justino, técnica do NGACB, por toda a ajuda na realização dos procedimentos laboratoriais deste trabalho. Aos colegas desbravadores da turma de doutorado 2009 do PPGBAN, por compartilharmos a insegurança, mas termos a coragem para sermos pioneiros. Aos professores doutores Albert David Ditchfield, Alfredo Langguth, Leandro Rabello Monteiro, Leonora Pires Costa, Marcos Vinícius Carneiro Vital e Sérgio Lucena Mendes, que aceitaram o convite para participar na banca de examinadora da tese e, certamente, contribuirão substancialmente para o aprimoramento desta pesquisa. iii Aos colegas do Laboratório de Mastozoologia e Biogeografia da UFES, por toda ajuda durante a realização do trabalho. Agradeço em especial à Ana Carolina Loss, pela enorme ajuda nos trabalhos no NGACB e pela cessão de sequências de DNA. Aos curadores e técnicos responsáveis pelas coleções examinadas, pela ajuda durante as visitas: Adriano Paglia e Raquel Moura (UFMG), Helio de Queiroz Boudet Fernandes (MBML), Márcia Jardim, Daniela Sanfelice e Glayson Bencke (MCNFZB), Teresa Cristina Castelano Margarido e Gilda Maria Siqueira Tebet (MHNCI), Mario de Vivo e Juliana Barros (MZUSP), João Alves de Oliveira, Stella Franco e Sérgio Maia Vaz (MNRJ), Suely Marques Aguiar e Ana Cristina Braga dos Santos (MPEG), Mariluce Rezende Messias e Ravena Mendonça (UFRO). Aos pesquisadores Antônio Rossano Mandes Pontes e Cláudia Regina Silva, pela cessão de alíquotas de tecido. Às pessoas que me auxiliaram em minhas visitas às coleções durante o doutorado. Ao meu amigo Israel de Souza Pinto, por me hospedar no Rio de Janeiro. Às minhas amigas Bárbara Maria Andrade Costa e Daniela Rossoni, por me hospedarem em São Paulo. Ao Instituo Federal de Educação, Ciência e Tecnologia do Espírito Santo, pelo apoio e concessão de horário especial de trabalho que possibilitaram a realização deste trabalho, e aos colegas professores e técnicos administrativos que souberam compreender certas necessidades durante meu doutorado. A Coordenação de Aperfeiçoamento de Pessoal de Nível Superior do Ministério da Educação (Capes/MEC), pela concessão de recurso financeiro (PROAP) utilizado para a realização da algumas visitas a coleções. Ao Programa Taxonomia do Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), pelo apoio financeiro que viabilizou boa parte das visitas às coleções. A todos que de alguma forma contribuíram para a realização desse trabalho, muito obrigado. iv Sumário Lista de figuras: ..................................................................................................................... vi Lista de tabelas: ................................................................................................................... viii 1. Resumo .............................................................................................................................. 1 2. Abstract .............................................................................................................................. 3 3. Apresentação ...................................................................................................................... 5 4. Capítulo 1- Filogenia molecular e divergência morfológica de ouriços-cacheiros (Rodentia: Erethizontidae) ................................................................................................... 10 4.1. Resumo .................................................................................................................... 11 4.2. Introdução ................................................................................................................ 12 4.3. Materiais e Métodos ................................................................................................. 14 4.3.1. Exemplares examinados ................................................................................... 14 4.3.2. Sequenciamento de DNA e inferência da filogenia ......................................... 16 4.3.3. Análises morfológicas e morfométricas ........................................................... 18 4.4. Resultados ................................................................................................................ 20 4.5. Discussão ................................................................................................................. 25 4.6. Referências Bibliográficas ....................................................................................... 32 4.8. Figuras ...................................................................................................................... 48 4.9. Tabelas ..................................................................................................................... 57 5. Capítulo 2 - Evolução fenotípica em ouriços-cacheiros (Rodentia: Erethizontidae) ....... 66 5.1. Resumo .................................................................................................................... 67 5.2. Introdução ................................................................................................................ 68 5.3. Resultados ................................................................................................................ 71 5.3.1 Desenvolvimento pós-natal ............................................................................... 71 5.3.2. Mapeamento de caracteres fenotípicos: ........................................................... 73 v 5.3.3. Processos evolutivos ........................................................................................ 75 5.4. Discussão ................................................................................................................. 77 5.4.1. Evolução da pelagem ....................................................................................... 77 5.4.2. Evolução craniana ............................................................................................ 81 5.4.3. Tendências evolutivas ...................................................................................... 83 5.5. Conclusão ................................................................................................................. 84 5.6. Metodologia ............................................................................................................. 85 5.6.1. Análise de caracteres fenotípicos e hipótese filogenética ................................ 85 5.6.2. Mapeamento e análise da evolução dos caracteres morfológicos .................... 88 5.7. Referências Bibliográficas ....................................................................................... 90 5.8. Anexo 1 .................................................................................................................... 98 5.9. Figuras ...................................................................................................................... 99 5.10. Tabelas ................................................................................................................. 118 6. Conclusão ....................................................................................................................... 121 vi Lista de figuras: Capítulo 1: Figura 1. Filogenia de Erethizontidae inferida a partir da matriz com sequências dos genes Cit b e vWF utilizando máxima parcimônia, máxima verossimilhança e inferência bayesiana .............................................................................................................................
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